View source for Biology:Zupaysaurus

{{Short description|Extinct genus of dinosaurs}}
{{Speciesbox
| fossil_range = Norian<br />~{{fossil range|221|206}}
| image = Zupaysaurus skull photo.png
| image_caption = The skull of ''Zupaysaurus''
| display_parents = 2
| taxon = Zupaysaurus rougieri 
| parent_authority = Arcucci & Coria 2003
| authority = Arcucci & Coria 2003
}}

'''''Zupaysaurus''''' ({{IPAc-en|ˌ|z|uː|p|eɪ|ˈ|s|ɔː|r|ə|s}}; "ZOO-pay-SAWR-us") is an extinct [[Biology:Genus|genus]] of early theropod [[Biology:Dinosaur|dinosaur]] living during the Norian stage of the [[Earth:Late Triassic|Late Triassic]] in what is now {{wipe|Argentina}}. Fossils of the dinosaur were found in the [[Earth:Los Colorados Formation|Los Colorados Formation]] of the Ischigualasto-Villa Unión Basin in northwestern Argentina. Although a full skeleton has not yet been discovered, ''Zupaysaurus'' can be considered a bipedal predator, up to {{convert|4|m|ft}} long. It may have had two parallel crests running the length of its snout.
 
== Discovery ==
Discovered in May 1997 by Santiago Reuil ("Vultur"), part of the crew of Guillermo Rougier, it was later described by Arcucci and Coria and published in 2003. The name ''Zupaysaurus'' is composed of the Quechua word ''supay'' meaning "devil" and the [[Social:Greek language|Greek]] word ''{{lang|grc-Latn|sauros}}'' ({{lang|grc|σαυρος}}) meaning "lizard"; thus "devil lizard". In Incan mythology, ''[[Religion:Supay|supay]]'' was both the god of death and ruler of the ''[[Unsolved:Pacha (Inca mythology)#Uku Pacha|ukhu pacha]]'', the Incan underworld. The [[Biology:Type species|type species]] was named ''Z. rougieri'' in the honor of Guillermo Rougier, the scientist who led the expedition which discovered and collected the [[Biology:Holotype|holotype]] (original specimen) PULR-076. ''Zupaysaurus'' was first described and named in the scientific journal ''Ameghiniana'' by {{wipe|Argentina}} paleontologists Andrea Arcucci and Rodolfo Coria in 2003.<ref name="arcuccicoria03"/>

== Description ==
[[File:Zupaysaurus skull.jpg|thumb|left|Skull diagram]]
''Zupaysaurus'' was a medium-sized theropod. An adult [[Biology:Skull|skull]], measured approximately {{Convert|450|mm|in|0|abbr=on}} in length, suggesting a body length of approximately {{Convert|4|m|ft|0|abbr=on}} from its snout to the tip of its tail. In 2010 [[Biography:Gregory S. Paul|Gregory S. Paul]] gave a length of 6 meters (20&nbsp;ft) and a weight of 250&nbsp;kg (550&nbsp;lbs).<ref>{{Cite book|last=Paul|first=Gregory S.|title=The Princeton Field Guide to Dinosaurs|url=https://archive.org/details/princetonfieldgu00paul_514|url-access=limited|publisher=Princeton University Press|year=2010|location=New Jersey|pages=[https://archive.org/details/princetonfieldgu00paul_514/page/n75 75]}}</ref> In 2016 it was given a much smaller size of 4.2 meters (14&nbsp;ft) and 70&nbsp;kg (154&nbsp;lbs).<ref>{{Cite book|last=Molina-Pérez & Larramendi|title=Récords y curiosidades de los dinosaurios Terópodos y otros dinosauromorfos|publisher=Larousse|year=2016|location=Spain|pages=253}}</ref> Like all theropods, ''Zupaysaurus'' walked only on its hindlegs, leaving its forelimbs free to grasp its prey. The length of the neck bones recovered suggests that this genus has a rather long neck. Like the coelophysoids, ''Zupaysaurus'' has a kink in its snout, between the premaxillary and [[Biology:Maxilla|maxilla]]ry bones of the upper jaw. It is estimated that ''Zupaysaurus'' had 24 teeth and an intermandibular hinge is present in the lower jaw.<ref name="arcuccicoria03">Arcucci, A.B. & Coria R.A. 2003. A new Triassic carnivorous dinosaur from Argentina. ''Ameghiniana'' 40(2):217-228.</ref> Only one specimen of ''Zupaysaurus'' is known to science. The [[Biology:Holotype|holotype]] specimen was designated PULR-076, which consists of a nearly complete skull which was very well preserved, the right shoulder girdle, the lower right leg and ankle, and twelve [[Biology:Vertebra|vertebra]]e from the neck, back, and hips. Additional material of a smaller individual found at the same site may or may not belong to ''Zupaysaurus''.
As ''Zupaysaurus'' was originally described, the head bore two thin parallel crests on top of the skull, similar to theropods like ''[[Biology:Dilophosaurus|Dilophosaurus]]'' and ''[[Biology:Coelophysis kayentakatae|Coelophysis kayentakatae]]''. These crests are thought to have been formed by the [[Biology:Nasal bone|nasal]] bones solely, unlike those of many other theropods which also incorporated the [[Biology:Lacrimal bone|lacrimal]] bones. Crests on the skull were pervasive among theropods and may have been used for [[Biology:Animal communication|communicative purposes]] such as species or gender recognition.<ref name="curriezhao93">Currie, P.J. & Zhao X. 1993. A new carnosaur (Dinosauria, Theropoda) from the Jurassic of Xinjiang, People's Republic of China. ''Canadian Journal of Earth Sciences'' 30: 2037-2081.</ref> However, more recent analysis of the skull has cast doubt on the presence of these crests in ''Zupaysaurus''. An unpublished abstract presented at a recent conference indicated the structures initially identified as crests were in fact the lacrimal bones displaced upwards during the process of [[Biology:Fossil|fossil]]ization.<ref name="ezcurranovas05">Ezcurra, M.D. & Novas, F.E. 2005. Phylogenetic relationships of the Triassic theropod ''Zupaysaurus rougieri'' from NW Argentina. Presented in August 2005 during the [http://acd.ufrj.br/mndgp/2clpv/en/index.html II Latin American Congress of Vertebrate Paleontology] {{webarchive |url=https://web.archive.org/web/20060504212911/http://acd.ufrj.br/mndgp/2clpv/en/index.html |date=May 4, 2006 }} in Rio de Janeiro, {{wipe|Brazil}}. This analysis will be published in peer-reviewed print form later in 2006. A summary of the talk can be seen [http://dml.cmnh.org/2005Aug/msg00175.html here].</ref> Other cranial ornamentation included a rugose laterally-projecting lacrimal ridge on the top of the skull.
[[File:Zupaysaurus.jpg|thumb|Life restoration of ''Zupaysaurus'' in a resting pose]]
A diagnosis is a statement of the anatomical features of an organism (or group) that collectively distinguish it from all other organisms. Some, but not all, of the features in a diagnosis are also autapomorphies. An autapomorphy is a distinctive anatomical feature that is unique to a given organism or group. According to Ezcurra (2006) and Ezcurra and Novas (2006), ''Zupaysaurus'' can be distinguished based on the following characteristics: the maxillary fenestra is within the [[Biology:Antorbital fenestra|antorbital fossa]] (according to Ezcurra, 2006), the rostral process of the lacrimal is ventrally bowed (according to Ezcurra, 2006), the ventral process of the squamosal is kinked (according to Ezcurra, 2006), wide contact between squamosal and quadratojugal (according to Ezcurra, 2006). The maxillary-jugal ventral margin describes an obtuse angle in lateral view (according to Ezcurra and Novas, 2006), a notch on the dorsal margin of the ascending process of the maxilla, relating to horizontal ramus of the lacrimal is rostrally tapering onto the forked caudal tip of the ascending process of the [[Biology:Maxilla|maxilla]]; (according to Ezcurra and Novas, 2006), a [[Biology:Lacrimal bone#In dinosaurs|lacrimal]] with a highly pneumatized antorbital recess (according to Ezcurra and Novas, 2006), a short and square-shaped retroarticular process of the mandible (according to Ezcurra and Novas, 2006), the [[Biology:Cnemial crest|cnemial crest]] is poorly developed (according to Ezcurra and Novas, 2006).

== Classification ==
''Zupaysaurus'' was classified as the earliest known tetanuran theropod due to several features of its skull, dentition, and hindlimb. However, several features typical of more [[Biology:Basal (phylogenetics)|basal]] theropods were also noted by the original authors.<ref name="arcuccicoria03"/> Analyses by Carano (2005), Tykoski (2005), and Ezcurra and Novas (2005) have classified ''Zupaysaurus'' as a coelophysoid related to ''[[Biology:Segisaurus|Segisaurus]]'' and probably ''[[Biology:Liliensternus|Liliensternus]]'', though more basal than ''[[Biology:Coelophysis|Coelophysis]]''.<ref name="ezcurranovas05"/><ref name="carranoetal05">Carrano, M.T., Hutchinson, J.R., & Sampson, S.D. 2005. New information on ''Segisaurus halli'', a small theropod dinosaur from the Early Jurassic of Arizona. ''Journal of Vertebrate Paleontology'' 25(4): 835-849.</ref><ref>Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Dissertation. University of Texas at Austin. 553 pp.</ref> Yates (2006) found ''Zupaysaurus'' to form a group with ''Dilophosaurus'' and ''[[Biology:Dracovenator|Dracovenator]]'', placing it in a monophyletic [[Biology:Dilophosauridae|Dilophosauridae]].<ref name="yates06">Yates, A.M., 2006 (for 2005). "A new theropod dinosaur from the Early Jurassic of South Africa and its implications for the early evolution of theropods." ''Palaeontologia Africana'', '''41''': 105-122.</ref> But later studies found ''Zupaysaurus'' to be a sister taxon sister to a clade containing [[Biology:Dilophosauridae|dilophosaurids]], ceratosaurs and tetanurans.<ref>Smith N.D., Makovicky P. J., Hammer W. R. & Currie P. J. 2007 Osteology of Cryolophosaurus ellioti (Dinosauria: Theropoda) from the Early Jurassic of Antarctica and implications for early theropod evolution. Zool. J. Linn. Soc. 151, 377–421.</ref><ref>Nesbitt, S. J., Smith, N. D., Irmis, R. B., Turner, A. H., Downs, A., & M. A. Norell. 2009. A complete skeleton of a Late Triassic saurischian and the early evolution of dinosaurs. Science 326:1530-1533.</ref>

Below is a [[Biology:Cladogram|cladogram]] based on the phylogenetic analysis conducted by Sues et al. in 2011, showing the relationships of ''Zupaysaurus'':<ref name=sues11>Hans-Dieter Sues, Sterling J. Nesbitt, David S. Berman and Amy C. Henrici (2011). "A late-surviving basal theropod dinosaur from the latest Triassic of North America". Proceedings of the Royal Society B 278 (1723): 3459–3464</ref>

{{clade| style=font-size:85%;line-height:75%
|label1=[[Biology:Theropoda|Theropoda]]&nbsp;   
|1={{clade
   |label1=&nbsp;[[Biology:Herrerasauridae|Herrerasauridae]]&nbsp;
   |1={{clade
      |1=''[[Biology:Staurikosaurus|Staurikosaurus]]''
      |2={{clade
         |1=''[[Biology:Herrerasaurus|Herrerasaurus]]''
         |2=''[[Biology:Chindesaurus|Chindesaurus]]'' }} }}
   |2={{clade
      |1=''[[Biology:Eoraptor|Eoraptor]]''
      |2={{clade
         |1=''[[Biology:Daemonosaurus|Daemonosaurus]]''
         |2={{clade
            |1=''Tawa''
            |label2=&nbsp;[[Biology:Neotheropoda|Neotheropoda]]&nbsp;
            |2={{clade
               |label1=&nbsp;[[Biology:Coelophysidae|Coelophysidae]]&nbsp;
               |1={{clade
                  |1=''[[Biology:Coelophysis|Coelophysis]]''}}
               |2={{clade
                  |1=''[[Biology:Liliensternus|Liliensternus]]''
                  |2={{clade
                     |1='''''Zupaysaurus'''''
                     |2={{clade
                        |1=''[[Biology:Cryolophosaurus|Cryolophosaurus]]''
                        |2={{clade
                           |1=''[[Biology:Dilophosaurus|Dilophosaurus]]''
                           |2=[[Earth:Jurassic|Jurassic]] theropods (which include ceratosaurs and tetanurans)}} }} }} }} }} }} }} }} }} }}

== Paleoecology ==
''Zupaysaurus'' was discovered in red siliciclastic sediments at the "Quebrada de los Jachaleros" locality within the [[Earth:Los Colorados Formation|Los Colorados Formation]] of the La Rioja province in Argentina. This [[Earth:Geological formation|formation]] has been shown by magnetostratigraphy to date to the Norian stage of the [[Earth:Late Triassic|Late Triassic]] period, approximately 228 to 208 million years ago.<ref name="weishampeletal04">Weishampel, D.B., Barrett, P.M., Coria, R.A., Le Loueff, J., Xu X., Zhao X., Sahni, A., Gomani, E.M.P., & Noto, C.R. 2004. Dinosaur distribution. In: Weishampel, D.B., Dodson, P., & Osmólska, H. (Eds.). ''The Dinosauria'' (2nd Edition). Berkeley: University of California Press. Pp. 517-606.</ref> but has also been assigned to the slightly younger Rhaetian stage, which was approximately 208 to 201 million years ago.<ref name="heckertlucas98">Heckert, A.B. & Lucas, S.G. 1998. Global correlation of the Triassic theropod record. ''Gaia'' 15: 63-74. [not printed until 2000]</ref> Both specimens assigned to this genus are housed in the collection of the National University of La Rioja in La Rioja, Argentina.<ref name="arcuccicoria03"/>

The [[Earth:Los Colorados Formation|Los Colorados Formation]] was interpreted as an ancient floodplain and it was home to several types of early sauropodomorph dinosaurs (including ''[[Biology:Riojasaurus|Riojasaurus]]'', ''[[Biology:Coloradisaurus|Coloradisaurus]]'' , and ''[[Biology:Lessemsaurus|Lessemsaurus]]''), all of which shared the same [[Earth:Natural environment|paleoenvironment]] with ''Zupaysaurus''.<ref name="weishampeletal04"/> It is recognized as one of the earliest known faunal assemblages dominated by dinosaurs, which were 43% of the number of tetrapod species currently known. The non-dinosaurs that inhabited this locality included pseudosuchians, therapsids like Cynodontia, other early reptiles, and possible archosaurs.

== References ==
{{Reflist}}

== External links ==
* ''[https://web.archive.org/web/20130504013424/http://archosaur.us/theropoddatabase/Coelophysoidea.htm#Zupaysaurusrougieri Zupaysaurus]'' entry in The Theropod Database.

{{Taxonbar|from=Q131786}}

[[Category:Dinosaur genera]]
[[Category:Norian life]]
[[Category:Los Colorados Formation]]

{{Sourceattribution|Zupaysaurus}}